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PSYCHE

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Louis M. Roth.
The Male Genitalia of Blattaria. V. Epilampra spp. (Blaberidae: Epilamprinae).
Psyche 77:436-486, 1970.

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THE MALE GENITALIA OF BLATTARIA. V.
EPILAMPRA SPP.
(BLABERIDAE : EPILAMPRINAE) .
BY Louis M. ROTH
Pioneering Research Laboratory
U. S. Army Natick Laboratories
Natick, Massachusetts 01 760
'The genus Epilampra is one of those assemblages which have de- veloped within the tropics of both hemispheres a vast number of species, often quite distinct, again closely related and difficult to distinguish.
With a general type of coloration the fluctuations of which make definite and exact characterization difficult, if not at times virtually impossible, it combines a uniformity of development in numerous other features, that in general in the family are suffi- ciently varied to prove of value to the systematic student. To add to the uncertainty of a situation difficult at best, we find many of the published descriptions almost valueless to aid in the recognition of these forms. As a whole the genus is one of the most difficult, ob- scure and generally unsatisfactory to study in the entire Blattidae." (Rehn and Hebard, 1927, p. 209).
Princis (1967) lists 60 species of Epilampra all of which are found only in the New World. At least five of these species [atriventris (Saussure) , cribrosa ( Burrneister) , ferruginea ( Brunner) , proximo (Brunner) , and verticalis (Burmeister) 1 have males with tergal glands, and their genitalia are so distinctly different from the males which lack tergal glands that I
(1970) have placed them in the
genus Poeciloderrhis Stil. This study of about 30 of the remaining 55 species of Epilampra listed by Princis ( 1967) shows that the male genitalia are useful not only for specific determinations of many species, but they may also indicate species relationships. MATERIALS AND METHODS
The technique of preparing slides of genitalia has been described in earlier papers (Roth, 1969b, 1970).
The source of each of the specimens illustrated is given, using the following abbreviations: (AMNH) = American Museum of Nat- ural History, New York; (ANSP) = Academy of Natural Sciences, Philadelphia; (BMNH) = British Museum (Natural History), London; ('CUZM) = Copenhagen University, Zoological Museum, Denmark; (L) = Zoological Institute, Lund, Sweden; (MCZ) =



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Fig. I, Male genitalia (dorsal view) of Epilampra abdomenninpurn from Tapurucuara, Rio Negro, Brazil. (C = cleft of Ll; LIZ first sclerite of left phallmnere; Lib =: setal brush of LI; L3vm ==: median sclerite L2 ventromedial) ; L2d = dorsal sclerim of 12; P = prepuce; R2 :^ hooked sderite of tight phallomere).
Museum of Comparative Zoology, Harvard University, Cambridge, Mass.; (N) = U. S. Army Labs., Natick, Mass.; (USNM) = United States National Museum, Washington, D.C. Geographical collection data, if known, follow these abbreviations. The number preceding the abbreviations refers to the number assigned the speci- men and its corresponding genitalia (on a slide) which were deposited in the museum indicated. These numbers are used in the text where the identifications of certain species are discussed. If known, the taxonomists who identified the species are given. In several cases these specialists of the Elattaria disagreed in their determinations, emphasizing the difficulty in identifying species of *!era from literature descriptions. Unfortunately male type tna- terial was not always available so that several questions still remain unresolved. In spite of this drawback the results point up the value of using male genitalia in the taxonomy of a difficult genus. The phallorneres characteristic of Ep2ampra male genitalia are shown in Figure I.
Prepuce - Usually distinctively shaped with a definite marginal outline and often covered by microtrichia (Fig. I, P) .



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438 Psyche [December
Lad - This sclerite (Fig. I, L2d) is always separated from L2vm (Fig. I) by a membrane, and may or may not be clearly sep- arated from and lie above the prepuce. In some species (e.g. Mexi- cans Group) most of Lad is a sclerotized and integral part of the prepuce.
R2 (retractable genital hook, Fig. I, R2) -A subapical incision is often found on the ventral surface (Fig. 18, SI) . LI -A deep lateral curved incision or cleft is present (Fig. I, C) and a setal brush (Fig. I, Lib) may or may not be found on the dorsal surface.
Rehn and Hebard ( 1927, p. 210) without specifying the characters used, tentatively erected 5 species Groups for Epilampra, primarily for West Indian species. These Groups and the species included were : I. Cubensis Group : - cubensis Bolivar.
2. Mexicana Group: - insularis Bolivar, tainana Rehn and Hebard.
3. Abdomennigrum Group : - abdomennigm (De Geer), mona Rehn and Hebard.
4. Burmeisteri Group : - gundlachi Rehn and Hebard, burmeis- teri (Guirin) , wheeleri Rehn, haitensis Rehn and Hebard, sabulosa Walker.
5. Grisea Group : - quisqueiana Rehn and Hebard. Rehn and Hebard (1927) indicated that other species would prob- ably fall into these groups. The male genitalia do not support the placement of a number of the above species in the groups erected by Rehn and Hebard. Based on genital characters I suggest the follow- ing species groups of Epilampra: Mexicana, A bdomennigrum, Bur- moisten, Sodalis, Shelfordi, Heusseriana, and Yersiniana. Mexicana Group
[Epilampra mexicana Saussure (Figs. 2-13) ; E. fallax Saussure and Zehntner (Figs. 14-23) ; E. conferta Walker (Figs. 24-43)]. This group includes species in which most of Lad is a flattened sclerotized plate which is an integral part of, and does not lie above, the prepuce (Fig. 5). In mexicana only a small part of L2d on the left side is separated from and lies above the prepuce, and on the right side the L2d tapers and extends upward toward the Lzvm (Figs. 2, 5, 8, 10, I I, 12). In fallax the lateral extension of the right side of Lad is quite long (Figs. 14, 17, 210, 21). In some con- ferta the L2d extends well beyond the left side of the prepuce (Figs. 24, 27, 29-32, 34) and the extension on the right may vary consid- erably in length. The prepuce of mexicana is deeply notched and is readily distinguished from the other members of the group.



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Roth - Blattaria
Figs. 2-1 3. Cockroach male genitalia. Epdampra m~xiruna. 2-4. (63 USNM). Guatemala. 5-7. (109 USNM). Turrialba, Coma Rica. 8-9. (106 USNM). Turrialba. Costa Rica- 10. (11 1 USNM). Chis. Soyalo, [Rt. 195, Km 241. Mexico. (2-10, det. Gurney). 11. (52 AMSP). Central Mexico. 12-13. (110 USNM). Ver. Rio TacoSopan, [Rt. 195, Krn 241, Mexico. (11-13 det. Roth). (L2d = dorsal acleritc of L2; P = preptice). (scale = 0.2 mm)




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440 Psyche [December
Figs. 14-23, Cockroach male genitalia, Epitampra jailax. 14-16. ( 51 ANSI'). Sapucay, Paraguay (Jet. Roth), 17-18. (66 USNM). Santa Cata- rina, Brazil (det. Albuquerque), (SI ^= Subapical Incision). 19-20. (73 WSNM).
Santa Catarina, Brazil. (det. Albuquerque). 23-23. (103 USNM). Rio Lujer, Buenos Aires, Argentina (det. Albuquerque). (in Fig. 21 the prepuce is collapsed), (scale "= 0.2 mm)



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Figs. 24-35, Cockroach male genitalia. Epiiumpra ronfcrtu. 24-Zb. ( 15+ US). El Valle, Panama (det. Princis). 27-28. (136 USNM). Barro Colorado Uland, Panama. (det. Princis). 29. (135 USNM). Barro Col~radO Island, Panama. (det. Roth). 30. (50 ANSP). ChirKpi, Panama. (labeled sligmosa in pencil), 31. (105 USNM). San Isidrodel, General Cattago, Costa Rica (det. Princis). 32-33. (HZ USNM). Napo, Santa Cecilia, Rio A , Ecuador. (det. Roth). 34-35, (133 USNM). Same locality as 32-33. (det. Roth). (scale == 0.2 mm)




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Psyche [December
i
s
, ,
t
b
1"
Figs. 36-46.
Cockroach male genitalia. 36-38. (4 CUZM). E. conferta. Callanga, Peru. (det. Princis) . 39-40. (121 USNM) . E. cozferta. Callanga, Dept. of Cudco, Peru. (dct Roth). 41-43. (67 USNM). Epilampra ~p. Rio Blanco or Ecuador. (det as E. mexicam by Albuquerqut). The sctal brush 5-1 Fig. 43 is very lightly pigmented and difficult to see in the photograph. 44-46. (168 USNM). Eftihmpra sp, Cundinamarca, Colombia. (wale = 0.3 mm)




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19701 Roth - Blattaria 443
In species of the Mexicana Group, R2 (Figs. 3, 6, 15,18, 22, 25, 28, 33, 37, 40, 42, 45) has a subapical incision, and a setal brush (Figs. 4, 7, 9, 13, 16, 19, 23, 26, 35, 38, 43, 46) occurs on LI. Princis ( I 958, p. 63) synonymized Epilampra stigmosa Giglio-TOS with Epilampra conferta Walker.
The specimens determined by
Princis (Figs. 24-28, 31) as conferta are similar to a specimen, in the Philadelphia Academy collection, which was labeled (in pencil) E. stignzosa (Fig. 30). However, the E. conferta identified byHebard, Albuquerque, and Rehn (Figs. 229-237) have entirely different geni- talia from Princis' conferta (Figs. 24-28, 31, 36-38). The type of Walker's conferta (Brazil) is a female, whereas Giglio-Tos' type of stigmosa is a male.
According to Gurney (personal communication) "E. stigmosa G.- T. was based on 4 males from Ecuador. Giglio-Tos also described E. josephi from 2 males from S. Jose, Ecuador (stigmosa was from the valley of Santiago, Ecuador) ; they apparently were similar to stigmosa for the most part. . . . No. 76 [conferta, det. Albuquerque] (Figs. 235-237), seems rather small to be conferta, judging from Walker's description (though type is a female), but size may be quite variable. I would be inclined to use the name stigmosa for No. 132 [Figs. 32-33], No. 134 [Figs. 24-26], and No. 136 [Figs. 27-28], and perhaps No. 105 [Fig. 311, but am more uncertain about No. 76 [Figs. 235-2371 being conferta. . . . No. 105 [Fig. 311 has the face darker than 132, et. al., the interocular space is wider, and the ventral surface of the abdomen is much darker; however, if the genitalia agree this may be just variation." It is apparent that Gurney is not convinced that stigmosa and con- ferta are the same species. However, for the present, I am following Princis' conclusions. It is highly probable that more than one spe- cies is involved here which are very similar in external appearance. The problem may be partly solved by examining the male genitalia of the type of stigmosa. Unfortunately the Type of conferta, as in- dicated above, is a female. The prepuce and L2d of conferta speci- mens from Ecuador (Figs. 32, 34) differ somewhat from these structures in specimens from Panama and Costa Rica (Figs. 24, 27, 29, 30, 3 I ) and from specimens from Peru (Figs. 36, 39). The R2
(Figs. 37, 40) of Peruvian males are noticeably stouter than the genital hooks (Figs. 25, 28) of Panamanian specimens. Epilam- pra conferta may well be a complex of sibling species. The genitalia of two undetermined or questionably determined species belonging to this group are shown in Figures 41-46. One
specimen from Ecuador, determined by Albuquerque as E. mexicana



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444 Psyche [December
is not this species, according to Princis (personal communication) and its genitalia (Figs. 41-43) are distinctly different from those of mexicana (cf. Figs. 2-13).
A bdomennigrum Group
[Epihzpra abdomennigrum (De Geer) (Figs. 50-55) ; E. maya Rehn (Figs. 47-49) ; E. sagit& Hebard (Figs. 59-67) ; E. taira Hebard (Figs. 56-58) ; E. grisea (De Geer) (Figs. 68-96) ; E. jorgenseni (Rehn)
(Figs. 97-1 13) ; E. berlandi Hebard (Figs. I 14- 117); E. guianae Hebard (Figs. 119-127)]. In this group L2d is a thick, black, variably shaped sclerite, which lies above the prepuce.
The presence of a setal brush on LI dis- tinguishes it from the following Burmeisteri Group. The size of the setal brush is inter- and intraspecifically variable and sometimes the area covered by the setae is small, or the setae are lightly sclerotized (Figs. 58, 99) and difficult to see. The hook (R2) has a subapical incision in all the species listed in this Group Epilampra maya (Figs. 47-49), until recently considered a synonym of E. abdomennigrum (Figs. 50-55), was shown to be a valid species by Roth and Gurney (1969). They illustrated the genitalia of a large number of individuals of both species to show the extent of variation which occurs in the phallomeres. Epilampra abdomenni- grum is found in South America, Trinidad, and the West Indian Islands of Grenada and St. Lucia, but whether it occurred in Puerto Rico was uncertain (Roth and Gurney, 1969). The Puerto Rican record reported here (Figs. 50-52) suggests that Rehn and Hebard (1927, p. 228)
were probably correct in regarding Sein's ( 1923) record of wheeleri in Puerto Rico as actually being abdomennigrum. Epilampra maya occurs in Central America and Mexico. The male
taken in Boston Quarantine
(Figs. 47-49) had Jamaica as the lo-
cality but it is possible that the specimen boarded ship in a Central American port.
Rehn (1902) stated that E. maya is closely related to E. conspersa and E. azteca and that it is separated from the latter by the shape of the supraanal plate. E. maya is very close to abdomennigrum with which it has been confused', and the genitalia of azteca (Figs. 247-249) are decidedly different and I have placed it in the Bur- rneisteri Group.
Hebard (1929, p. 366) stated that E. sagitta is near (by markings) E. colunzbiana and E. opaca. However, the shape of L2d and prepuce of s&ta (Figs. 59, 62, 65) appear to be closer to those of abdomen- nigfium (Figs. 50, 53-55) than to columbiana (Figs. 208-219) and



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Psyche [December
Figs. 56-67. Cockroach male genitalia. 56-58. (83 USNM) . Epilum$ra taira. Surinam (dm. Gurney) (arrow in 58 indicates setai brush). 59- 67, Efilampra sagitta. 59-61. (74 U-SNM). Amapii, Brazil (det. Albuqucr- que ; confirmed by Frincis), 62-64. (182 ANSP). Type 1135. Teffe, Ama- zonan, Brazil. 65-67. (N). Florea, Manaun, Brazil (det Hoth). (scale = 0.2 mm)




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19701 Roth - Blattaria
447
fig^. 68-76. Cockroach male genitalia. EpiSaiafra &#a. 68-70. (42 ANSP) . Bariica, British Guiana (diet. Hebard). 71-73. (34- AMNH) . Surinam (labeled E. lu(i/upa Rehn, a synonym of grisea). 74-76. (157 USNM). Rosario, Lake Rogagua, Bolivia (det. Roth). (scale = 0.2 mm)



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Psyche [December
Figs. 77-96. Cockroach male genitalia. Epil~mpra grisea. 77. (.W USNM). Trinidad, British West Indies. 78. (93 USNM). Lelydorp, Suma- trawcg, Surinam. 79. (94 USNM). Surinam. 80. (96 USNM). Popogai- mama Creek, Surinam. 81-82. (156 USNM) . Blancaflor, Beni, Bolivia. S3-93. (N). 83-85. Flores, Manaua, Brazil. 86. Adolpho Ducke Forestry Preserve, about 60 Km. from Manaua, Brazil. 87. Puraquequara, Rio Negro, Amazonan, Brazil. 88-89. Moura, Rio Negro, Amazonas, Brazil. 90-93. Tapurucuara, Rio Negro, Amazonas, Brazil. 94. (107 USNM). 95. (108 USNM). At Plant Quarantine, Miami, Florida; with some bromeliads, possibly from Peru. 96. (65 WSNM).
At Miami, in plane, probably from
South America. (all det. Roth). (scale = 0.2 rnm]



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19701 Roth - Blattarh 449
Figs. 97-108.
Cockroach male genitalia. EfUainfra jorgensmi. 97-105. Farawpea of Efilamfra dgmaiiphra Rehn (=: E. jorqenseni). Miaionea, Argentina. 97-99. (86 AMP).
(arrow in 100 indicates setal brush}. 100- 102. (96 ANSP). 103-105. (48 AMSF). (Fig. 103 is a ventral view; part of the prepuce (arrow) in this specimen is missing). 106-109. (164 USNM), Between Coronel Oveido and Asuncion, Paraguay (det. Roth). (scale = 0.2 mm)




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450 Psyche [December
Figs. 109-117. Cockroach male genitalia. 109-1 13. Epitampra jargmseni. 109-1 10. (94 ANSP). Paratype of Ep'itampra stigm.uii$hora Rdn (= E. ]or- genscd). Misiono, Argentina. 111. (95 ANSP). Misiones, Argentina (det. Rehn). 112-113. (81 ANSP). same data as figs. 109-110. 114-117. Epilanpra
bcrhdi. 114, (47 ANSP}. Provincia Sara, Bolivia (det. Hebard). 115- 117. (83 ANSP). Paratype. Icafio, Santiago del Estero, Argentina. (arrow in 117 indicates setal brush). (scale = 0.2 rnm)



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Roth - Btattaria
Figs. 118-126. Cockroach male genitalia. Ep lampra guianae, 118-120. (35 ANSP), Paratype, Rockstone, British Guiana. 121-323. (91 USNM). Surinam. 124. (90 USNM). 125. (89 USNM). Brokopondo, Surinam. 126. (92 USNM). Surinam, (rips of prepuce (arrows) in figureh 125 and 126 missing). ( 121-126, det. Roth). (scale == 0.2 mm)



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452 Psyche [December
Figs. 127-135. Cockroach male genitalia. 127-133. Epilampra burmei- attri}. 127-129. (2.5 MCZ). Cuba (labeled E. raraibea S. and Z., which ia a synonym of bttrmeisteri). 130-131. (24 MCZ). Yimque de Baracoa, Oriente Province, Cuba (Set. Gurney). 132-133. (39 ANSP). Cuba (dct. Rehn). 134-135. (21 MCZ). Epilamfra takuna. Mountains north of Imias, Orierrte Province, Cuba (det. Gurney). The LI of taintma wag lost in preparation of the slide and the species is tentatively placed in die Bw- mei$lerf Group; all other Cuban Spilumpra belong to this Group. (scale = 0.2 mm)




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Figs. 136-144.
Cockroach male genitalia, Epiiumpra qtkqueiana. Para- types. 136-158. (37 ANSP). 139-141. (84 ASSP). San Lorenzo, Province of Saman4, Dominican Republic, Hispaniola. 142-144. (82 ANSP). San Francisco Mountains, Province of Santo Domingo, Dominican Republic. (scale = 0.2 mm)




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454 Psyche [December
opaca (Figs. 178-195 ) which I place in the Burmeisteri Group be- cause of the absence of a setal brush on LI. The prepuce of E. taira (Fig. 56) is relatively broad and extends well beyond the margins of the L2d. These structures resemble those of E. opaca but the right side of the prepuce of opaca is more extended and pointed (Fig. I 78).
The prepuce of E. grisea is greatly elongated, tapers to a point posteriorly, and shows remarkably little variation (Figs. 68, 71, 74, 77-81, 83-92, 94-96) over a wide geographical range. Based on Princis' (1967) catalogue, new geographical records for this species are Bolivia (Figs. 74-76)', Trinidad (Fig. 77)) and possibly Peru (Fig. 95).
The L2d of E. jorgenseni is unusual in being very large and deeply indented. The left side of the L2d tends to follow the contour of the underlying prepuce (Figs. 97, 100, 103, 106, 109, 112). One of the specimens (det. Rehn) apparently has an aberrant shaped L2d ( Fig. I I I ) . The genitalia of E. berlandi (Figs. I I 5-1 I 7 ) are very similar to those of jorgenseni and support Hebard's ( 1921, p. 283) claim that the two species are closely related. The Paraguayan record for jorgenseni (Figs. 106-108) is new.
The prepuce of E. guianae is unique for this group, and i,s deeply notched (Figs. I I 8, 12 I, 124-126) ; in this respect the prepuce is reminiscent of that of mexicana (Figs. 2, 5, 8, 10-12)) though the general shape is different between these 2 species. The prepuce of one unusual specimen of guianae has 2 indentations in the prepuce (Figs. 125).
Burmeisteri Group
[Epilampra burmeisteri (Guirin) (Figs. I 27- I 33 ) ; E. tainana Rehn and Hebard (Figs. 134-135) ; E. quisqueiana Rehn (Figs. I 36-144) ; E. sabulosa Walker (Figs. 145-150) ; E. wheeler! Rehn (Figs. 15 I- 156) ; E. gundlachi Rehn and Hebard (Figs. 157-162) ; E. haitensis Rehn and Hebard (Figs. 163-165) ; E. hamiltoni (Rehn) (Figs. 166- 168) ; E. bromel'iadarum (Caudell) (Figs. 169-1 7 I ) ; E. exploratrix (Gurney)
(Figs. 359-361 ) ; E. gatunae (Hebard) (Figs. 172-174) ; E. fugax (Bonfils) (Figs. I 75-177) ; E. opaca (Walker) (Figs. 178- 195 ) ; E. substrigata Walker (Figs. 196-207) ; E. columbiana Saus- sure (Figs. 208-219) ; E. latifrons Saussure and Zehntner (Figs. 299- 301) ; E. histriga Walker (Figs. 220-228) ; E. thunbergi Princis (Figs. 238-243) ; E. castanea Brunner (Figs. 244-246) ; E. azteca Saussure (Figs. 247-280) ; E. crossea Saussure (Figs. 293-298) 1. This Group is essentially similar to the Abdomennigrum Group



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19701 Roth - Blattaria 45 5
but is separated from it by the absence of a setal brush on LI. R2
usually has a subapical incision but a few species lack this character. The prepuce is usually well defined but in a few species it is markedly reduced.
The Burmeisteri Group includes the largest number of species of Epilampra and may be further divided into subgroups based on the relative sizes, shapes, and extent of development of L2d and the prepuce. Although not all of the species will fit readily into the fol- lowing subgroups many of them do show a close relationship and I believe an attempt at sub-divisions is worthwhile. Subgroup A:
(burmeisteri, tainana, quisqueiana, sabulosa, wheel- eri, gundlachi, haitensis, hamiltoni, bromeliadarum, gatunae, f ugax) : In thi,s subgroup the area of Lad is relatively small in relation to, and covers only a small anterior part of the prepuce. In burmeisteri (Figs. 127, 130, 132), quisqueiana (Figs. 136, 139, 142), and sabulosa (Figs. 145, 148, 150) the L2d is roughly divided in 2 parts, the left half usually being larger and sometimes more darkly pigmented than the right half. Rehn and Hebard (1927, p. 233) compared quisqueiana with grisea and substrigata. The prepuce and L2d of quisqueiana are closest to sabulosa and differ noticeably from those of substrigata (Subgroup B, Figs. 196, 199, 202-203, 205-206) and grisea (Abdomennigrum Group, Figs. 68, 71, 74). Epilampra gundlachi has been confused with burmeisteri (Rehn and Hebard, 1927, p. 223) but the male genitalia of these 2 species are distinctive (cf. Figs. 157-159 and 127-129). According to Rehn and Hebard (1927, p. 228), the nearest relative of haitensis is wheet- en. However, the Lad of haitensis (Fig. 163) is closer to gundlachi (Figs. 157, 160) than it is to wheeleri (Figs. 151, 154) ; note the pointed spur on the left side of L2d in gundlachi and haitensis, and its absence in wheeleri.
For almost 50 years the status of the genus Audreia Shelford has been in question. Hebard (1920, p. 92-93) stated "First steps only have as yet been taken to separate properly the generic units related to EpiZampra, in which partial to complete reduction in organs of flight has occurred. At the present time, as was the case with Shel- ford in 1910, insufficient material is at hand to allow a proper revi- sion to be made. A number of species are before us which must be assigned to this genus as characterized by Shelford, but which indicate the presence of at least four distinct groups, though females alone of the majority of species are represented. Larger series and male examples will be needed before it can be determined whether these



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456 Psyche
[December
Figs. 145-156. Cockroach mate genitalia, 145-150. EfHamPru tubula~a, 145147. (33 AMNH). La Moriniere, Haiti (det Rchn). 148-149. (51 ANSF). Bourdon, Haiti (det Rehn). 150. (149 ANSP). Porto Prince, Haiti. 151-156. Kpilumpra wheeleri. 151-153. (32 AMNH). Adjuntas, Porto Rico (det. Kehn; from specimen shown in figure S in Rehn and Hebard, 1927). 154-156. (102 USNM). El Yunque, Porto Rico (det. Roth). (scale : 0.2 mm)




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Figs. 157-165.
Cockroach male genitalia. 157-162. Epiiurnpm gmdlachi. 137-159. (97 USKM). Cayamas, Cuba (det. Hebard), 160-162. (23 MCZ).
Mountains north of Imias, Oriente Province, Cuba (det. Gurney). 163- 165. (69 USNM). Epiiampra haitensis, Port-au-Prince, Haiti (det. Albuquer- que). (scale = 0.2 mm)




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458 Psyche [December
Figs. 166-177. Cockroach male genitalia. 166-16S. (175 USNM). Epi- lampra hamiitmi. Pico, Turquino, Cuba (det. Gurney). 169-171. Epilampra bromel'adarum. Panama. ((let. Huber) . 172-174. (184 USNM) . Epilampra gahinae. Pozo Azul, San Joab, Costa Rica (det. Gurney). 175-177. Ep'lamfra fu@ux. Paratype of hdreia fugax Bonfils. Saint Francoise Anse-a I'Eau, Guadeloupe. (scale = 0.2 mm}




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Figs. 178-186. Cockroach male genitalia. Epiiutnpra apaca, 178-180. (3% ANSP). St. Jean du Maioni, French Guiana (det. Hebard). 181-153. ( 17 CUZM ). Essequibo, British Guiana (det. Princis). 184-186. (80 USNM). Arnapi, Brazil (det, a'i near licrlundi by Albuquerque, and as sag.'tta by Princis'). (scale = 0.2 mm)




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460 Psyche
[December
Figs. 187-195. Cockroach male genitalia. Efilamfra upam. 187-189. (70 USNM). Amapi, Brazil (det. as near bdandi by Albuquerque). 190, (SE USNM). Brownsberg, Wakti-basoe Creek, Surinam (det. as sag tfu by Frin- cis). 191-193. (125, 127, and 126 USNM). Napa Santa Cecilia, Rio Aguarico, Ecuador. 194-195. (89 and 87 ANSP). Provincia Sara, Dept. Santa Cruz, Bolivia, (all det. Roth). (scale = 0.2 mm)



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Figs. 196-207,
Cockroach male genitalia. E$i[umpra d&gata. 196-19s. (129 USNM).
Nap0 Santa Cecilia, Rio Aguarico, Ecuador (det. Roth). 199-200. (46 ANSP) . Vi!lavicensio, Co~ombia (det Heba td) . 201-202. (6s WSNM), Anabern, Colombia (det. Ahquerqw]. 203-204. (71 USNM). Territ. Amazonas, San Fdo. Atabapo, Venezuela (det. Albuquerque). 205. (128 USNM). Napo Santa Cecilia, Rio Aguarico, Ecuador. (det. Princis). 206-207. (79 USNM). Territ. ArnapA, Brazil (det. Albuquerque). (scale = 0.2 mm}




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462 Psyche
[December
Figs, 208-219, Cockroach male genitalia. E#itumpra coiumbiunu. 20% 210. (49 ANSF). 211-213. (138 GSNM). Barro Colorado Island3 Panama ((let. Roth). 214-216. (98 WSNM). Sierra Campana, Panama (det. Gurney). 217-218- (137 USNM). Barro Colorado Island, Panama (det. Roth). 219. (154 USXM). La Carnpana, Panama (det, Roth). (wale = 0.2 mm)



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Roth - Blattarh 463
Figs. 220-22s.
Cockroach male genitalia. Epi/ampra bmisiriga, 220-222- (78 USNM). Piracicaba, S3o Paulo, Brazil (det. Gurney), 223-225. (104 USNM). Brazil (det, A[buquerque). 226, (114 USNM), South of S50 Paula, Brazil. 227. (90 ANSP). Guaraja, Sgo Paulo, Brazil (det. hy Hebard as E. ddicd~ Hebard, a 3ynonym of busistriga). 22K (77 USNM). Same
locality as figures 220-222; L2d is turned on its side (det. Roth). (scalt = 0.2 mm)




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464 Psyche [December
groups represent distinct genera, or are simply striking divisions in the genus Audreia, comparable to the many striking divisions in the genus Epi1amjv-a as at present defined. . . . Whether the tegmina and wings are reduced, truncate, lateral and lobiform, or absent, are features which in the Blattidae are known to be often utterly worth- less from a generic standpoint. . . . JVithout the sexes of each species, we do not feel in a position to characterize Audreia more definitely than has been done in the meager description given by Shelford." A few years later Rehn and Hebard (1927, p. 204) commented further on the genus Audreia stating that it ". . . is composed of a small number of species described from the tropics and subtropics of both hemispheres, the majority, however, tropical American. The species much resemble certain forms referred to the genus Epilampra, but a11 possess reduced or subquadrate tegmina in both sexes, these subquadrate or distally emarginate in most of the forms . . . Until the genus EpiZampra as a whole is critically studied, and our knowl- edge of the extent to which brachypterism occurs in that assemblage is more complete, it is unwise to do other than follow Shelford's use of the generic name Audreia. We can say, however, that the genus CaZoZampra, to which a number of the species now placed in the more recently described Audreia were originally referred, is well distinct from Audreia of Shelford. The West Indies possess two species which can logically be assigned to Audreia, one from higher mountains of eastern 'Cuba, the other from Blue Mountains of Jamaica. The possibility that these may be members of an ancient relic fauna forces itself upon one, although the converse argument that tegminal reduction has been brought about by adjustment to a peculiar and restricted montane environment cannot be ignored." The male genitalia of CaZoZampra carinulata Saussure, the species which Hebard (1920, p. 92) selected as the type for the genus Audreia Shelford show 2 distinct differences from the genitalia of most species of Epilampra. The hooked right phallomere (Fig;. 348, 351) lacks a subapical incision and is relatively stout. The hook from the specimen shown in Fig. 348 tends to resemble the hook of E. sodalis (Fig. 309) but is shorter, The L2d of A. carinulata (Figs. 347, 350) is a flattened sclerotization of the preputial membrane; the remainder of the prepuce is shapeless. In most EpiZanzpra, the prepuce has a well defined shape and is densely covered by micro- trichia.. The LI of A. carinulata lacks a setal brush (Figs. 349, 352). The genitalia of 5 other species of Audreia (Figs. 166-177) differ from those of A. corinzdata, and are similar to the genitalia of certain other species of EpiZmzpra. Rehn and Hebard ( 1 ~ 2 ~ ~ p. 205) stated



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19701 Roth - Blatturiu 465
that Audreia haitziltoni ". . . is clearly congeneric with carinulaia (Saussure j of Central America, the genotype, as selected by Hebard, although it has a number of features of difference." However, the L2d and prepuce of hmdtoni (Fig. 166)) bronze~iadu~um (Fig. 1691, gatunae (Fig. 172)) fugax (Fig, 175)) and explorairix (Fig. 359) are so typical of E$iZampra that I assign them to this genus. The R2 of fugax (Fig. 176) lacks a subapical incision; this incision is present in the other 4 species of "Audreia" (Figs. 167, 170, 173, 360) but is much reduced in bro~nc~iadarzm (Fig, 170)~ gatunae
(Fig. I 73) ) and expZorutrix (Fig. 360). The LI'S (Figs. 168) 171, I 74, 177, 361) of all 5 species lack a setal brush as do the other members of the Burnzeisteri Group.
Princis (1967) lists g ,species of Audreia) two of which (A. cicatricosa Rehn, and A. jamaicanu Rehn and Hebard) I have not seen. It is possible that these species are also EpiZanzpr~, Princis in- cluded EpiZarnpra heusseriana under Audreia but its genitalia (Figs. 302-307) are basically those of Epifanzpru and I have placed it in a separate group (see below j. The male of Audreia catharina Shelford has tergal glands on tergites I and 2. Its genitalia are basically sim- ilar to species of "Epilmnpra" that possess tergal glands and I placed it in the genus Poeciloderrhis (Roth 1970). For the present I suggest that the genus A,udreia) as represented by carinulata, be retained until a more detailed study is made of other morphological characters of this species. Subgroup B :
(opaca, substrigata, colmnbiana, Zutifrons, basistriga, thunbergi, castunea) : The species in this subgroup have a relatively large L2d which overlies a considerable area of the prepuce. According to Princis (1958, p. 62) Walker's E. opaca is a syn- onym of his E. substrigata. Princis (personal communication) ex- amined the types of opaca and substrigata ". . . and could not find any noteworthy differences. Hebard had never seen the types and he thought them to be two different species. I supposed that Hebard's records from French Guiana [opaca] could be correct, whereas his record of substrigata from Colombia evidently relates to another species. This was, however, a pure speculation of mine.'' The geni- talia of Hebard's xu bstrigata from Colombia (Figs. I 99-200) clearly differ from those of specimens he considered to be opaca from French Guiana ( Figs. I 78- I 80) . Princis' su bstrigata ( Fig. 205 ) is similar to Hebard's substrigata specimens (Fig.
199). The genitalia of a
specimen from British Guiana at the CUZM, determined as opaca by Princis (probably before he considered it to be a synonym of substrigato) are similar ( Figs, I 8 I -I 83 ) to Hebard's opaca (Figs- I 78-180).




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Pry che
[December
Figs. 229-237. Cockroach male genitalia. Epilumpu sp. 229-231. (44- ANSP).
Provincia Sara, Dep. Santa Cruz, Bolivia (det. as E, cenferta by Hebard). 232-234. (43 ANSP). Parintins, Para, Brazil (det, as E. confertu by Rehn). 235-237. (76 USNM). Serra do Navio, Territ. Amapa, Brazil {det. E. conferta by Albuquerque; recorded in Albuquerque and Gurney, 1962, p. 242). (see discussion on page 443). (scale = 0.2 mm)



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19701 Roth - Haitaria 467
Figs. 23&246. Cockroach male genitalia. 238-240. (18 CUZM). Efi- lampra thuftbtfgi. Rio de Janeiro, Brazil (det. Princis). 241-243. (11 BMNH). E. thunbergi. Theresopolis, Santa Catarina, Brazil (det, Princh ; this specimen was determined as E. testatea Brunner by Hebard). 24't- 246. (18 BMNH).
EfiSamfra castmea (dct. Princia; this specimen was determined as "Epilamfra" functkallir Walker by Hanitsch). (scale = 0.2 mm)




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Psyche [December
Figa. 247-259. Cockroach male genitalia. Efi!amfra acteca. 247-249. (171 USNM) . Pdenque, Chiapaa, Mexico (det. Roth) . 250-252. (41 ANSP). Pozo Azui, Costa Rica (det. Hebard). 353-254. (141, 140 USNM). Barro Colorado Island, Panama (det. Roth). 255. (144- USW). Cerro Campana, Panama (det. Roth). 256, (20 CUZM). Taboga, Panama (det. Frincis). 257-259. (141, 142, 140 USNM). Barro Colorado Island, Panama (det. Roth). (scale = 0.2 mm)




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19701 Roth - Blatfaria 469
Figs. 360-271. Cockroach male genitalia. Epilamfra wteca. 260-262. (84 USNM). Wakti-basoe Creek, Brown~beig, Surinam. 263-264, <S6 USNM), Fhedra, Surinam. 265-266. (87 USNM), Same data a3 Figs. 260- 262 267-268. (85 USNM). Same data as Figs. 260-262. (260-268 det. Gurney). 269-271. (13 BMNH) .
Trinidad (very light specimen labeled
"extreme reccsfnve color").
(det, Hebard). (scale = 0.2 mm)




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470 Psyche [December
Figs. 272-280. Cockroach male genitalia. Efilumpra asteca. 272-274. (75 USNM). Territory Amazonas, Mt. Marahuaca, Venezuela (det. AI- buquetque). 275-277. (131 USNM) . Napo Saata Cecilia, Rio Aguarico, Ecuador (del Roth). 278-280. (I30 USNM). Same data as Figs. 275-277. (dm. Roth). (scale = 0.2 mm)




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19701 Roth - Blattaria 47 1
Gurney (personal communication) commented that "Judging from Hebard's descriptive remarks about substrigata Walker, he, Albuquer- que, and Princis apparently have correctly identified it, but unfor- tunately there is no type locality for it, and the types of both it and opaca Walker are females . . . On the other hand, the type locality of opaca is Demerara, British Guiana, so Hebard may have had it from his French Guiana collections."
For the present I am considering opaca and substrigata to be dis- tinct. In addition to the marked differences in L2d and prepuce of these 2 species, the hook (R2) of substrigata (Figs. 197, 200, 201, 204, 207) is usually distinctly wider than that of opaca (Figs. 179, 182, 185, 188). Hebard (1926, p. 201) stated that E. opaca . . . may prove to be a synonym of the Brazilian E. inaculicollis (Serville), and the Ecuadorean E. stigmosa Giglio-Tos may fall in the same synonymy. Numerous distinct, though easily confused, spe- cies are known to belong to this group and, until a better concept of the distribution and individual variation within these is formed, we believe it best to use the name opaca." According to Princis stigmosa is a synonym of E. conferta (see discussion under Mexicam Group). Hebard (1921, p. 136) stated that substrigata is closely related to p-isea ". . . though separable by numerous features." The genitalia of substrigata relate it more closely to cohmbiana (Figs. 208-219) and opaca (Figs. 1 78-186), than to grisea (Figs. 68-76). The difficulty in identifying some of these species is shown by the fact that two specimens (Figs. 184-186, I 87-189) determined as closely related to "herlandi" (cf. Figs. I I 5-1 I 7) by Albuquerque and Gurney ( 1962, p. 243) are similar to Hebard's opaca. Princis determined one of these specimens (Figs. 184-186) as well as one from Surinam (Fig. 190) as E. sagitta. Gurney examined these specimens and in the absence of a careful study of types and genitalia felt that external features suggested the occurrence of more than one species. There can be little doubt that the genitalia of the Type of E. sagitta (Figs. 62-64) and what is here considered to be opaca are distinctly different. In sagitta LI has a setal brush (Figs. 61, 64, 67) (Abd~fiienni~runz Group) and the tip of the prepuce is directed more posteriorly (Figs. 59, 62, 65). In opaca there is no setal brush on LI (Figs. 180, 183, 186, 189) (Burmeisteri Group) and the tip of the prepuce is directed laterally (Figs. 178, I 81, 184, 187, 190- 195 ) .
Hebard (1920, p. 98) stated that E. columbiana ". . . is extremely close to E. mexicana Saussure and may eventually prove to be a geo- graphic race of that insect." These are unquestionably distinct spe-



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472 Psyche
[December
Figa. 281-289.
Cockroach male genitalia. Ephpra ap, D. 281-283. (115 USNM). Turrialba, Costa Rica. 284-286. (133 USNM). Same data as Figs. 281-283 (281-286 (Set. as E. avteca by Princia). 287-289. (139 USNM). Cerro Carnpana, Panama (det. Roch). (scale = 0.2 mm)



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fly
Figs. 290-298, Cockroach male genitalia. 290-293. {17S USNM) . Epi- imnpra axteca. Holotype of E. coiorata R. S. Albuquerque and Gurney. Rio Fdicio, Amapi, Brazil. 293-298. Ep hpra mrssea. 293-29s. (40 ANSP). St. Jean du Maroni, French Guiana (det. Hebard). 296-298. (85 ANSF). Para, Brazil. (male = 0.2 mm)




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474 Psyche [December
Figs. 299-307.
Cockroach male genitalia. 299-301. (25 MCZ). Ep'lumpra latifrom Pernambuco, Brazil. (det. Rehn) . 302-3 07. Epilumpra heusseriafia. 352-304. (151 WSNM). Cerro Montevideo, (det Albuquerque). 305-307, (160 L'SNM). Rio Grande da SuI, Brazil (det Roth) (scale = 0.2 mm)



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19701 Roth - Blattaria 475
Figs. 308-319,
Cockroach male genitalia. EjnJampra sodalls. (all Para- types of Epilamfra cinnamomea Hebard. St. Jean da Maconi, French Guiana. 30B-310. (36 ANSP), 311-313. (93 ANSP). 3M-316. (80 ANSP). 317-W. (91 ANSP). (scale = 0.2 mm)




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Psyche [December
Figs, 320-32%. Cockroach male genitalia. 320-32s. Epilampra mduli~. 320-322. (92 ANSP),
St. Jean du Maroni, French Guiana (Para-e of E. cinnumam~u Hebard], 323-325. (48 USNM). Surinam (det. Roth) . 326- 328. (8% AMP). Park Brazil (det. Rehn; arrow in Fig. 327 point^ to a thin line representing fie mbapical iuci9ion). (3cale = 0.2 mm)



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19701 Roth - Blattaria 47 7
cies and their genitalia are so different that by my criteria I place them in different groups. The shape of the L2d of E. columbiuna ( Figs. 208, 2 I I, 2 14, 2 I 7, 2 I 9) is distinctly fishlike and it is easily distinguished from mexicana (Figs. 2, 5, 8, 10, I I, 12). The L2d and prepuce of E. Zatifrons (Fig. 299) are very si~nilar to those of abdomennigrunz (Figs. 50, 53-55) ; the LI of Zutifrons lacks a setal brush (Fig. 3011, whereas these setae are present in abdomennigrum (Fig. 52).
The unique tarsal-clawlike shape of the L2d of basistriga (Figs. 220, 223, 226-228) distinguishes this species from any other Epi- Zampra. Hebard (1929, p. 369) believed that E. deZicata (Fig. 227) (= basistrigu) seemed to be near E. berlandi, and E. jorgenseni and apparently even more closely related to Audreia catharina; this con- clusion is not supported by genitalia.
E. berlandi (Figs. I 14-1 17)
and jorgenseni
(Figs. 97- I I 3) are in the A bdo?nennigrunz Group ; and Shelford's Audreia catharina with genitalia (Figs. 37-39 in Roth 1970) completely different from those of EpiZampra is now in the genus PoeciZoderrhis.
Two species, E. thunbergi (Figs. 238-2431 and E, castanea (Figs. 244-2461 are apparently closely related; the R2's (Figs. 239, 242, 245) and LI'S (Figs. 240, 243, 246) are especially similar. Hebard apparently had misidentified one of these specimens (Figs. 241-243) of thunbergi as E. testacea. Princis ( 1949, p. 65) discussed thunbergi (Type: c? ?, without abdomen) and compared it with grisea and also stated that E. substrigata IValker may prove to be a synonym of thunbcrgi. The genitalia of grisea (Figs. 68-96) and substrigata (Figs. 196-207 ) are distinctly different from Princis' thunbergi (Figs. 238-240). Hebard (1929, p. 365) stated that what he considered to be testacea [= thunbergi] (Figs. 241-2431 was closely related to guianae (Figs. I I 8- I 26) ; but this conclusion is not supported by the genitalia wlich are markedly different.
The specimen of E. castanea was recorded by Hanitsch (1931, p. 385) as EpiZmzpra puncticoZZis. E. fiuncticoZ/is is now in Rhab- doblatta, a genus not found in the New IVosld (Princis, 1967). This specimen is from the Fry Collection in BMNH and Hanitsch in reporting it commented that "No particulars are available concern- ing the specimens from the late Xlr. Alexander Fry's collection. Some of the material seems &lalayan, but the rest is almost certainly Neotropical." Fry lived at one time in Rio de Janeiro and this specimen most probably came from there.
Subgroup 'C (azteca, crossea) : The prepuce is usually poorly de- veloped or indefinitely outlined; if the prepuce is clearly outlined, its surface sculpturing is scalelike.




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478 Psyche
[December
The genitalia of E. azteca from different localities suggest that two species and possibly three may be included in this taxon. In specimens from blexico (Fig. 2471, Costa Rica (Fig. 250)) and Panama (Figs. 253-256)) L2d is irregular in outline and lies above the prepuce; the prepuce is irregular in outline and some portions tend to blend into the surrounding membrane. The L2d in specimens from Trinidad (Fig. 269)) Surinam (Figs. 260) 263, 265) 267)) Venezuela (Fig. 272)) and Ecuador (Figs. 275) 278)) is knoblike in shape and appeal-s to be an outgrowth of the prepuce. The L2d in two specimens from Costa Rica (Figs. 281, 284) (det. a; azteca by Princis), and one from Panama (Fig. 287) differs dis- tinctly fsom the azteca just discussed; the prepuce is more clearly defined and its scalelike surface sculpturing also differs from the other azteca. This is probably an undescribed species and I am tenta- tively calling it Epilampra sp. D. Gurney (personal communication) has commented on E. sp. D. and states ". . . they (Nos. I 13, I 15, 139) are very much like azteca from Central America (Nos. 140, I 41, I 43, I 44) but differ in face markings, However, No. I 3 I from Ecuador is darker in general) has darker and larger face markings, and the ventral surface of the abdomen is mostly blackish) unlike I 15 et al. The type of aztcca is a male from 1\4exico because) though Cuba and Mexico were both mentioned originally, Cuba was elim- inated as type consideration by comments in the Biologia. TVe have a male from "Mexico" which in face markings is more like the Princis det. specimen than like 140, et. al.) so perhaps Pi-incis is right. Con- sulting the tj~pe of azteca should solve the problem.'' According to Albuquerque and Gurney ( 1962, p. 244)) E. colorata is related to the '(7nacuZifrons" Stil group. The genitalia (Figs. 290- 292) of the Type of colorata are very similar to those of E. azteca from 'Central America (e.g. Figs. 250-252). Gurney (personal com- munication) 1-e-examined the Type and stated that colorata is quite likely a syno,nym of azt~ca. The Type is smaller than many azteca but probably within the normal size range. TVith Gurney)s con- currence I consider colorata a sj~nonym of uzteca. The prepuce of E. crosseu (Figs. 293, 296) has no distinctive shape and is simply a membrane covered with microtrichia. Heusseriana Group
[E. heusseriana Saussure (Figs. 302-3071 ] At present, only E. hemseriana belongs to this group. The L2d
(Figs. 302, 305) of heusseriana is unusually large in relation to the prepuce. R2 has a subapical incision (Figs. 303) 306) and LI lacks a setal brush (Figs. 304, 307).




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19701 Roth - Blattaria 479
The battleaxe-shaped L2d is continuous with a sclei-otized portion of the prepuce and is not separated from the prepuce by a thin clear membrane (as is usual in the Abdomennigrum and Burmeisteri Groups). In the Mexicana Group) L2d is not a distinct sclerite lying above the psepuce but is a flat sclerotization lying on the same plane as the prepuce.
There has been some question about the placement of EpiZampra heusse~i~na Saussure, According to Hebard ( I 92 I ) this species ". . . has been assigned to the genera Calolampra and Audreia) due mainly to the fact that the type female had short truncate tegmina. Though this is true for the female sex) the male before us is seen to have fully developed organs of flight, and shows no characters which would warsant its being placed other than in the genus Epilmzpra." Princis ( I 967) lists heusseriana under Audreia apparently following Albuquerque ( 1964). The male genitalia of heusscrima are typical of many other Epilampra and tend to support Hebard's placement of the species.
Sodalis Gi-oup
[Epilampra sodalis JValker (Figs. 308-3281 ] In this group which includes E. sodalis and possibly several unde- termined species) there is no distinctive prepuce and L2d is a single more 01- less clawlike sclerotization. In a recent paper I indicated that the male of E. sodalis had tergal modifications on segments 3 and 4 (Roth) 1969a, p. 202) Table 10) footnote b). I have examined males of E. cinnmzomea Hebard, which Princis synonymized with sodalis Walker (Princis) 1958, p. 16)) and found that the males lack tergal glands. I also examined Walker's Type of sodalis and con- cur with Princis that cinnamomea is a synonym of this species. The male genitalia of sodalis (Figs. 308-328) are distinctive; L2d re- sembles a single tassal claw, and R2 is rather stout with a subapical "incision" represented by a fine line which is best seen in a lightly sclerotized specimen (Fig. 32'7) arrow). The genitalia ( Figs. 329-33 I ) of the specimen previously reported by me as sodalis (Roth, 1g6ga) (here designated as E~ilampra sp. A) are only subtely different from those of sodalis; L2d (Fig. 329) and R2 (Fig. 330)
are slightly more slender in sp. A. However, there is a striking difference in the pronotal markings of these two species. In sodalis the microspots are all small and more or less the same size whereas in sp* A there are distinctly large spots) inter- spersed among small ones. These two forms are probably sibling species. Species A is the only Epilampra I know in which male tergal



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Psyche [December
Figs, 329-337. Cockroach male genZta1ia. 329-331. (182 USNM). F#i- fampra sp. A. Tapurucuara, Rio Negrox Brad, 332-337'. Epilampra sp. C* 332-334. (152 USNM) . Camp Rangel* Arague, Vemzuela. 335-336. (55 USNM). Same data as Fip. 332-334. 337. (112 USNM), AT, Rancho Grandc, Venezuela.
(scale = 0.2 mm)




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19701 Roth - Bluttadu 48 1
Figs. 338-146. Cockroach male genitalia. 338-343. Epilamfira sp. 5. 338-340. (10 BMNH). Fancina, Vera Paz. 341-343. (72 USNM). Territi Amazonaa, Mt. Marahuaca, Venezuela (det a3 E. sodah by Albuquerque). 344-346. (153 USNM). Efilampra sp. (near sp, R). Taken at Hoboken Quarantine (Venezuela?). (scale = 0.2 mm)



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482 Psyche [December
Figs. 347-358. Cockroach male genitalia. 347-352. A̤drei rarkulata (Saussure). 347-349. (176 USNM). La Palma, Costa Kica. (det. Rehn). 350-352. Volcan Barba, Costa Rica, (det. Fisk). 353-355. ( USNM). Efilampfa yhiana. SSo Paulo Rio Cubatso, Brazil (det Princin). 356- 358. (181 ANSP). EpHampra shflfordi. Type 5345. El Coredo, Cauca, Colombia. (scale = 0.2 turn)




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19701 Roth - Bhttiaria
483
Figs. 359-361.
( 108 MCZ). Cockroach male genitalia of EpUuntpra ex- plorutrix (Gurney). Holotype 25559 of Asdreia exploratrix Gurney. BUEDOS Aires, Trinidad Mts,, Santa Ciara Province, Cuba. (the left side of L2d
[Fig. 359, arrow] was torn due to pressure of the coverslip, and the tip of R2 [Fig. 360, arrow] was accidentally cut off). (scale =. 0.2 mrn) modifications are found on segments 3 and 4; the genitalia do not conform with those of Poecihderrhls whose species (formerly in Epilainpra) have tergal glands on segments I and 2 (Roth, 1970). Two or three other species may belong to. this group. Epilmnpra sp. B.
(Figs. 338-3431 has a very small clawlike Lad (Figs. 338, 341) and its R2 (Figs. 339, 342) differs markedly from soil/ilis (cia Fig. 327) ; Albuquerque misidentified this species ( Figs. 34 I -343) as soda&. Princis determined one of these specimens as 1;. fiihtmh'intta but its genitalia (Figs. 338-340) are quite different from specimens which I and Gurney (personal coimnunication ) conshlri- to be columbianu (cf. Figs. 208-2 19). Another specimen, Ep'ilftmpra sp, is very near sp. B.
(Figs. 344-346) ; it has an Lad (Fig. 344) slightly different in shape from the other two specimens :md its V.2 (Fig. 345) is more elongate. In Epilampra sp. C (Figs. 32-~7), Lad
(Figs. 332, 333) is not heavily sclerotized but is a liulloiv, membranous, pointed extension of the preputial membrane :md is covered by mkrotrichia, Its R2 (Figs. 333, 336, 337) is noticeably shorter than that of sp. B (cf. Figs, 339, 342). Yehiam Group
[EpHampra yersiriiana Saussure (Figs. 353-355) ] In E. yersiniuna the prepuce, when flattened, extends obliquely to the right of Lad (Fig. 333). The hook (R2) lacks a subapical in-



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484 Psyche [December
cision and its tip is nipple shaped (Fig. 354). The setal brush of LI extends vertically behind the darkly sclesotized cleft region (Fig. 355 1.
Though originally described as an Epilampra, y ersiniana has been placed in the genus Hedaia (by Saussure, Finot, Kirby, Hebard, and Rocha e Silva Albuquerque) and was listed by Kirby as a Rhabdo- blatta ( Princis, 1967) . Princis ( 1967) listed the species under Epi~ lanzpra and stated (personal communication) that "Hedaia is a Malagassy genus and I do not believe that it occurs in South Amer- ica. Probably a new genus is required [for yersiniana]." Shelfordi Group
[Epilampra shelfordi Hebard ( Figs. 356-358 ) ] E. shelfordi is the only species belonging to this group. It is unique in that L2d is absent and the preputial membrane is in the form of a rounded hollow cylinder (Fig. 356) covered with microtrichia. The R2 lacks a subapical incision (Fig. 357). No setal brush is present on LI
(Fig. 358).
According to Hebard (1919, pp. 106-
IO~), E. shelfordi ". . . belongs to an apparently exclusively South American group of the genus . . . To this group belong E. conspersa and E. agathina, of which single specimens are at hand. More mate- rial may show these forms to be generically distinct.'' The male genitalia of species of Epilampra are not only useful for specific determinations but can also be used to show species relation- ships. Thirty-six species of this genus have been divided into the following Groups and Subgroups :
I. Mexicana Group : - inexicana, fallax, conferta. 2. Abdomennigmnz Group : - abdomennigrum, nzaya, sagitta, taira, grisea, jorgenseni, berlandi, guianae. 3. Burmeisteri Group
a.
Subgroup A : - burmeisteri, tainana, quisqueiana, sabu- losa, wheeleri gundlachi, haitensis, hanziltoni, bromeliadarum, gat- tunae, fugax, exploratrix.
b.
Subgroup B: - opaca, substrigata, colunzbiana, latifrons, basistriga, thunbergi, castanea.
c. Subgroup 'C : - azteca, crossed.
4. Heusseriana Group : - heusseriana.
5. Sodalis Group : - sodalis.
6. Yersiniana Group : - ~ersiniana.
7. Shelf ordi Group : - shelfordi.




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19701 Roth Blattaria 485
Genitalic differences indicate that E. opaca is a distinct species and not a synonym of E. substrigata.
The male genitalia of 5 species of Audreia (hamiltoni, bromelia- darum, exploratrix, gatunae, and fugax) are so characteristic of Epi- Za?n$ra that I have transferred them to this genus. Epilanzpra colorata is synonymized with E. azteca. ACKNOWLEDGEMENTS
I thank the following for the loan of museum material: Dr. M. G. Emsley and Dr. N. Jago, formerly with the Philadelphia Acad- emy of Natural Sciences, Dr. A. B. Gurney, U. S. National Museum, Washington, D.C., Dr. Karl Princis, Zoological Institution, Lund University, Sweden, Dr. Jerome G. Rozen, Jr., American Museum of Natural History, New York, Dr. S. L. Tuxen Zoological Mu- seum, Copenhagen, Dr. David R. Ragge, British Museum (Natural History), London, Dr. Ivan Huber, Dr. J. Bonfils, and Dr. P. R. Castillo.
In addition I collected several species of Epilampra during Phase C of the Alpha Helix expedition to the Amazon in 1967. I tha,nk the National Science Foundation for support on the ~hazon expedition under Grant NSF-GB-5916.
I am grateful to Dr. K. Princis and Dr. A. Gurney for deter- minations and Mr. Sam Cohen for taking the photographs. REFERENCES
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1967. Orthopterorum Catalogus. Pars. 11 - Blattariae: Subordo Epi- lamproidea. Fam. : Nyctiboridae, Epilampridae. 's-Gravenhage, pp. 616-710.
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ROTH, L. M.
1969a. The evolution of male tergal glands in the Blattaria. Ann. Ent- omol. Soc. Amer. 62: 176-208.
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